Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 August 2020 | 12(11): 16407–16423
ISSN 0974-7907 (Online) | ISSN
0974-7893 (Print)
doi: https://doi.org/10.11609/jott.6510.12.11.16407-16423
#6510 | Received 01 August 2020 |
Final received 09 August 2020 | Finally accepted 19 August 2020
Use of an embedded fruit by Nicobar Long-tailed
Macaque Macaca fascicularis
umbrosus: II. Demographic influences on choices
of coconuts Cocos nucifera and pattern of forays to palm plantations
Sayantan Das 1,
Rebekah C. David 2, Ashvita Anand 3 ,
Saurav Harikumar 4, Rubina Rajan 5 &
Mewa Singh 6
1,6 Biospychology laboratory, Vijnana Bhavan, Institute of Excellence, University of Mysore, Mansagangothri, Mysuru, Karnataka 570006, India.
1 Wildlife Information Liaison
Development, No. 12, Thiruvannamalai Nagar, Saravanampatti - Kalapatti Road, Saravanampatti, Coimbatore, Tamil Nadu 641035, India.
2 Centre for Wildlife Studies,
37/5, Yellappa Garden, Yellappa
Chetty Layout, Sivanchetti Gardens, Bengaluru,
Karnataka 560042, India.
3 Foundation for Ecological
Research, Advocacy and Learning, No. 170/3, Morattandi,
Auroville, Tamil Nadu 605501, India.
3 InSeason Fish, Tarapore Avenue, Harrington Road, Chetpet, Chennai, 600031, Tamil Nadu 600031, India.
4 CR205B Bioscience Building,
Biological Sciences, Department of Biological Sciences, Faculty of Science
& Engineering, Culloden Road, Macquarie University, Sydney 2109, Australia.
5 Amity Institute of Forestry and
Wildlife, Amity Road, Sector 125, Noida, Uttar Pradesh 201303, India.
6 Zoo Outreach Organization, No.
12, Thiruvannamalai Nagar, Saravanampatti
- Kalapatti Road, Saravanampatti,
Coimbatore,
Tamil Nadu 641035, India.
1 sayantaniiser@gmail.com,2 rebekahcdavid@gmail.com,3
ashvitaa95@gmail.com, 4 saurav.hari-kumar@students.mq.edu.au,
5 rubina1611@gmail.com,6 mewasinghltm@gmail.com
(corresponding author)
Editor: Anonymity requested. Date of publication: 26 August 2020
(online & print)
Citation: Das, S., R.C. David,
A. Anand, S. Harikumar, R. Rajan
& M. Singh (2020). Use of an embedded fruit by
Nicobar Long-tailed Macaque Macaca fascicularis umbrosus:
II. Demographic influences on choices of coconuts Cocos nucifera and pattern of
forays to palm plantations. Journal of Threatened Taxa 12(11): 16407–16423. https://doi.org/10.11609/jott.6510.12.11.16407-16423
Copyright: © Das et al. 2020. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: American
Society of Primatologists, World Wildlife Fund for
Nature-India, Inlaks India Foundation and Science and Engineering Research Board, Government of India.
Competing interests: The authors declare no competing interests.
Author details: SD manages the
Nicobar Project of the Biopsychology laboratory at the University of Mysore. SD
is interested in behavior, ecology and environmental
conservation. RCD is a fundraising coordinator at the Center
for Wildlife Studies, Bengaluru and is interested in a range of issues
including conservation of natural world, philosophy, feminism, human rights,
business and politics. AA is interested in animal behavior,
human-animal interactions and wildlife conservation. SH is pursuing a Master’s
degree in Conservation Biology and desires to undertake research in
conservation sciences. RR is also pursuing a Master’s degree in Wildlife
Science and is inclined towards issues of human-wildlife interactions, animal behavior and urban ecology. MS is an eminent wildlife
biologist and specializes in primatology, conservation biology and psychology.
His areas of research specialisations include ethology, ecology, anthropology,
wildlife conservation and genetics.
Author contributions: SD and MS designed
the study and its methodologies; SD, RCD, AA, SH and RR undertook field data
collection; SD organized and assimilated the data; SD performed the analyses;
SD prepared the figures; SD drafted the manuscript; MS and SH provided
editorial inputs to the manuscript.
Acknowledgements: We are grateful to
the principal chief conservator of forests (Wildlife), Andaman & Nicobar
Forest Department, Port Blair for permitting us to carry out the study (Permit
No. CWLW/WL/134/Vol.XII/435). We were privileged to receive necessary
logistical and administrative assistances throughout the course of the study
from the divisional forest officer, Nicobar Division, range forest officers of Kamorta and Katchal, and
assistant commissioners of Campbell Bay and Nancowrie.
Our deepest obligations and appreciation extends to all the research assistants
who have worked with the Nicobar Project in various capacities and were
involved in field data collection. Ms.
Monica Harpalani and Mr. Bodhisatwa
Chaudhuri of the research team assisted in organizing and sorting the field
data from Katchal and Great Nicobar, respectively. We are indebted to our field assistants at Katchal, Mr. Mahesh and Mr. Sebastin. Dr. Swetashree Kolay and Mr. Nisarg Desai advised us on statistical analyses and we
sincerely thank them for their benevolence.
SD received grant from the American Society of Primatologists, the Inlaks India Foundation, Mumbai and the World Wildlife
Fund, New Delhi which supported RCD, AA, SH and RR. MS receives SERB’s
Distinguished Fellowship Award (Number: SB/S9/YSCP/SERB-DF/2018(1)).
Abstract: Adaptive pressures of
human-induced rapid environmental changes and insular ecological
conditions have led to behavioral innovations among behaviorally flexible
nonhuman primates. Documenting long-term
responses of threatened populations is vital for our understanding of species
and location-specific adaptive capacities under fluctuating equilibrium. The Nicobar Long-tailed Macaque Macaca fascicularis umbrosus, an
insular sub-species uses coconuts Cocos nucifera, an embedded cultivar
as a food resource and is speculated to have enhanced its dependence as a
result of anthropogenic and environmental alterations. We explored demographic patterns of use and
abandonment of different phenophases of fresh
coconuts. To study crop foraging
strategies, we recorded daily entry and duration of forays into coconut
plantations. We divided age-classes into
early juvenile (13–36 months), late juvenile (37–72 months), and adults (>72
months) and classified phenophase of coconuts into
six types. Consistent with the theory of
life history strategies, late juveniles were found to use a greater number of
coconuts, which was considerably higher in an urban troop but marginally higher
in a forest-plantation dwelling group.
Except in late juveniles, males consumed a higher number of coconuts than
females in the remaining age-classes.
Owing to developmental constraints, juveniles of both types used higher
proportion of immature coconuts though adults showed equitable distribution
across phenophases.
Pattern of entries to plantations and duration of forays were uniform
through the day in the urban troop but modulatory in the forest-plantation
group, perhaps due to frequent and hostile human interferences. Observations corroborating adaptations to
anthropogenic disturbances are described.
Keywords: Coconut phenophases, hard to process food, human-induced
rapid environmental change, human-macaque competition, dependence on coconut,
coconut-based resource competition, coconut consumption, Nicobar archipelago
Introduction
Among the many challenges that primates and their habitats face
globally, rapid and escalating anthropogenic changes in the age of the
Anthropocene are having an irreversible effect on primate populations leading
to exclusion, extinction (~60% of primate species, Estrada et al. 2017) and
severe constriction of ranges in most primate species (~75% of primate
species), (Estrada et al. 2017; Erinjery et al. 2017;
Kalbitzer & Chapman 2018). Although a few dietary and habitat generalist
primate species are beginning to show indication of behavioural adaptation to
anthropogenic habitats (McLennan et al. 2017; Santini et al. 2019), many
specialist primate species are trapped in their ecological niches constrained
by their phylogeny, life-history, physiology and/or limited phenotypic plasticity
(Vázquez & Simberloff 2002; Fisher & Owens
2004; Kalbitzer & Chapman 2018). Even among populations that are synanthropic/commensal to humans, many studies have
enunciated the impact of habitat modification on a variety of socioecological
(Back et al. 2019), parasitological (Kouassi et al.
2015; Zanzani et al. 2016; Kumar et al. 2018;) and
health variables (Kaur et al. 2008; Muehlenbein et
al. 2010). Many flexible populations of
Apes, Old and New world primate populations subsisting in anthropogenic
habitats especially of the genus, Pan, Macaca,
Papio, Cebus,
Cholorcebus, and Saimiri
exhibit evidences of compensating for dietary stress with expansion of dietary
resources (like crops and synthetic foods) and associated supplemental foraging
strategies (Pan, Hockings et al. 2015; Macaca,
Ilham et al. 2017; Brotcorne et al. 2017; Papio, Fehlmann et al.
2017; Cebus, Back et al. 2019; Cholorcebus, Thatcher et al. 2020; Samiri, Campêlo et al.
2019). Many of such food-enhanced
populations show complex sensorimotor intelligence associated with extraction
of embedded food resources and feed on food items novel to their ancestral diet
(e.g., oil-palm nut processing by Burmese Long-tailed Macaque, Luncz et al. 2017).
Alongside many novel frugivore-fruit relationships, the relationship
between the Nicobar Long-tailed Macaque Macaca
fascicularis umbrosus
(Images 1,2)and the coconut Cocos nucifera L., a perennial cash
crop is particularly intriguing since both the species have colonised the
Nicobar archipelago of the Andaman & Nicobar Islands. Although the nature of dependence of the
macaque species on wild varieties of coconuts occurring in the islands is
unknown, domesticated land races of coconuts have arrived on the island ~2,250
years ago (see Gunn et al. 2011; Niral & Jerard 2018). Groups
of macaques closest to coconut palm plantations are exposed to the drupe and
thus, familiar to ‘domesticated’ coconuts and coastal groups that have had
prolonged exposure to coconut palms have a much higher dependence than recently
exposed inland groups (Das et al. 2020).
Systematic destruction of habitats for expansion of coconut horticulture
and agriculture (Arora 2018), human habitations and defence establishments
along with environmental changes/catastrophes (aridity/Indian ocean tsunami)
have disproportionately affected groups on the edge of their habitats (Karnauskas et al. 2016; Reddy 2018), constituted largely by
coastal populations of long-tailed macaques (Umapathy
et al. 2003; Velankar et al. 2016). Under such circumstances, it becomes
essential to study the adaptive pressures of both, gradual and extreme habitat
alterations on coastal populations and the resultant behavioural responses,
especially in context of dietary expansion and foraging innovations. Since, many such dietary adaptations can have
adverse effects on survival and/or persistence of a species in an agriculture
ecotone especially, if these resources are shared or cultivated by humans
(Hockings et al. 2015; Hill 2017; Kalbitzer &
Chapman 2018), it becomes vital to study behavioural flexibilities to explicate
adaptive capacities of species and/or population(s) experiencing anthropogenic
pressures. Behavioural flexibilities
within a group, however, are not expressed identically across demographic
classes and age-sex class-specific strategies prevail as a result of distinct
life histories (Stamps & Krishnan 2017).
For instance, studying the dynamics of group fission in Sumatran
Long-tailed Monkey, van Schaik & Noordwijk (1985)
described age and social affiliation-specific disintegration of foraging
parties with large-bodied sub-adults foraging solitarily during fruiting
seasons. Even size and hardness of
fruits fed varied along the age-sex axes (van Schaik & Noordwijk
1985). Although many sub-species of
long-tailed macaques have been documented to feed on complex embedded resources
(like Opuntia spp., Tan et al. 2016; Terminalia catappa,
Falótico et al. 2017; Elaeis
guineensis, Proffitt et al. 2018) including usage
of stone tools to access few of them, variation in the use of these resource
items along demographic axes has been seldom investigated (c.f. intertidal
shellfishes, Gumert et al. 2011). We adopted the HIREC framework (human-induced
rapid environmental change) expounded by Sih et al.
(2016) to understand adaptive pressures specific to individual species along
with commensurate dietary flexibilities, adaptive potential, and overall
phenotypic flexibility in response to extreme anthropogenic changes to
ecosystems. We estimated that the
severity of HIREC would be compounded in an insular condition due to the
ecological fragility of island ecosystems leading to the exertion of stronger
adaptive pressures on coastal groups of long-tailed macaques than on
inland/mainland groups (e.g., many island populations of long-tailed macaques
(e.g., Malaivijitnond et al. 2007; Luncz et al. 2017) and capuchin monkeys show tool-use behavior (e.g., Barrett et al. 2018)). Despite phylogenetic
constraints on expression of behavior, we expected
insular populations of long-tailed macaques to express greater behavioral flexibility, quicker learning, proficient
extractive foraging and greater tendency of dietary expansion (e.g., Malaivijitnond et al. 2007; Tan et al. 2015, 2016). Thus, the human-macaque interface in the heterogeneous
habitat of Nicobar Islands creates a virtual experimental condition for
studying emergence of foraging and other dietary adaptations and/or innovations
under conditions of HIREC.
In the current study, we focused on how demographic categories, i.e.,
age and sex compared to each other and to other similar groups in their use of phenophase of coconuts.
We also aimed to study contingent acquisition and abandonment of
coconuts by age-classes and describe their probable causes. Based on the theory of life-history
strategies in macaques, we hypothesized that older juveniles (3–6 yrs) would feed on the highest number of coconuts followed
by adults (>6yrs) and younger juveniles (1–3 yrs)
due to the largest energy requirement of older juveniles among all
age-classes. Comparison of the two sexes
though is less straightforward since both, reproductive females and adult males
have high energetic requirements for procreation and for maintenance of larger
body size, respectively (e.g., Collins 1984; van Schaik & Noordwijk 1985).
Since procreation lasts for a shorter time scale than body maintenance,
we expected adult males to feed on a higher number of coconuts than adult
males. For the remaining age classes, we
expected no difference between the two sexes.
Because the husk and the shell of the coconut gets progressively tougher
and harder with development, we expected adults to process higher number of
mature coconuts than by juveniles though tender coconuts will continue to be
preferred choices by all age-classes due to the ease of extractive processing.
The marginal value theorem (MVT) within optimal foraging theory
postulates that the time spent in resource patches by individuals/groups
follows maximization of net energy, i.e., the difference in energy invested in
foraging and the energy gained by ingestion (Pyke et al. 1977; Charnov & Orians 2006). Group-level patterns of decisions pertaining
cultivar use and plantation visitation is comprehensively specified by MVT,
which assumes a greater prominence when conjoined to the HIREC framework since
cultivar (resource) attractiveness, cultivar (resource) value and risks from
human and non-human crop defenders are introduced as additional factors. In this study, we were interested in
expounding and contrasting patch entry and patch use by two groups with
different degrees of coconut-dependence, different experiences of human
hostilities and different distribution of coconuts, throughout the day. A secondary intent was to generate data that
would serve as a baseline for more detailed studies on movement and foraging
decisions in contested landscapes.
Further, we used the MVT framework within HIREC to obtain insights into
the processes governing entry/exit and patch usage dynamics of the focal
groups.
Methods
Study site
We undertook the study at Great Nicobar and Katchal
in the Nicobar archipelago of Andaman & Nicobar Islands lying between
93.634–93.953E & 6.735–7.229N, and 93.301–93.475E & 7.873–8.026N,
respectively (Figure 1). The major
forest types in these islands are the Andaman tropical evergreen forest and the
Andaman semi-evergreen forest (India State of Forest Report 2019). Due to their isolation from continental
mainland, the islands have high degree of endemism with an extremely poor
mammalian diversity (Nayar & Shastry
1987; Balakrishnan 1989; Rao 1989). The
Nicobar Long-tailed Macaque is found across all vegetation types in the
archipelago including littoral beach formations, mangrove vegetations on
coastal regions, low land swamps and inland wet evergreen vegetations (Hajra et al. 1999; Arora 2018). Over the past century, unregulated phases of
human migrations and unsustainable developmental initiatives have led to
large-scale deforestation on the eastern coast of the islands altering local climatic
conditions and threatening biodiversity.
Human settlements, agricultural/production landscapes and other
human-dominated spaces on the eastern coast are the primary centers
of human-macaque hostilities (Rajeshkumar 2017). We chose to study coastal groups of Nicobar
Long-tailed Macaque in the two islands that ranged within human-dominated
spaces and showed considerable dependence on anthropogenic food resources.
Study groups
We studied two groups of long-tailed macaques, one in each island. The study groups ranged in coastal areas of
the two islands. The first group, Temple
Run (TR) subsisted within a matrix of semi-urban area, patchily-distributed
native vegetation, advanced secondary forest and home garden/plantation of
Campbell Bay town in Great Nicobar.
Coconut palms occurred in sparse numbers within small (0.04ha) to
moderate-sized gardens (0.5ha) maintained at government offices, residential
areas, temples and other public spaces.
Therefore, TR had access to coconuts almost throughout the day (see Das
et al. 2020). The second group, Baywatch
(BW) used a cumulative coconut plantation area of 5.75ha spread across three
patches that ran along the northeastern coast of Katchal. Alongside,
the group also accessed semi-altered mixed evergreen forest and other coastal
native vegetations. The group had six of
(probably) seven–eight sleeping sites adjacent to palm plantations and largely
consumed coconut at dawn and dusk. For
information on the demographic structure of the two groups, see Das et al.
(2020). The troops used coconuts
considerably in general and used tougher and mature coconuts consistently, in
specific. Conclusively, the troops displayed remarkable proficiencies in
extractive foraging of coconuts signifying a long and an involved relationship
with the nut, however, both the troops faced immense hostility from humans/dogs
within agricultural and other anthropogenic landscapes as a result of crop
depredation. Even so, active dispelling
of macaques of both troops neither had an effect on their daily allocation of
time spent in coconut plantation nor on daily coconut consumption (Das et al.
2020).
Field methods
Post habituation of the two groups, we began data collection from March
2018 for a period of 24 months and 20 months for TR and for BW,
respectively. We followed TR from March
2018 to February 2020 and BW from March 2018 to October 2019. We divided the observation period into two
annual cycles which began in March and ended in February as a result of the
annual periodicity in coconut consumption (see Das et al. 2020). Due to our failure to identify immature
individuals of TR group in the first annual cycle of the study, we report the
results from the second annual cycle alone.
Groups were followed from dawn to dusk at least once a week and for a
minimum of five days in a month with sampling day considered as successful only
if all coconuts acquired by a troop were accounted for. We noted coconut acquisition by the groups
within an all occurrence behavioral sampling
framework with each session continuing for 10 minutes. We recorded entry and exit schedules into
coconut palm plantations of the troops, acquisitions of fresh coconuts from
direct (from palm) and indirect (from other individuals and from ground)
sources followed by their respective fates, i.e., either processed (if liquid
endosperm is accessed) or unprocessed (if liquid endosperm is not accessed) and
finally, age and sex classes of individuals (wherever possible) acquiring
them. The sampling challenges presented by
the two troops as a result of the habitats they occupied led to minor
difference in the field protocol followed.
This included an inability to record phenophases
of coconuts acquired by TR group as a result of inaccessibility to coconut
palms. For description of the six phenophases of coconut and their identifying features, see
Das et al. (2020). We combined the third
and the fourth phenophases due to similarities in
their developmental characteristics and for the purpose of easier
representation. Demography of the two
groups was assessed on a monthly basis.
Data analysis
We classified the life span of Nicobar Long-tailed Macaques into three
classes, 13–36 months (early juveniles, EJ), 37–72 months (late juveniles, LJ)
and >72 months (Adult, AD) based approximately on (1) coconut
handling/processing proficiency and on (2) conventional age classifications for
Macaques. We assessed the age-classes of
individuals on a monthly basis. For the
purpose of testing inter-annual consistencies, we partitioned the dataset of BW
into the two annual cycles described previously and presented data of TR over a
single annual period only. Whereas, to
contrast temporal visit patterns to palm plantations (within a day) by TR with
BW, we averaged data across the two annual cycles and represented them as
‘frequency of entry’ during 10 minutes slots along with corresponding time
spent in plantations.
Unprocessed coconuts emerge when macaques acquire coconuts directly
(from palm or ground) or indirectly (snatch from a conspecific) but leave them
unfed as a result of unsuitability of coconut (i.e., coconut is
diseased/disfigured/barren), incapability to process, mishandling (slippage
while on the palm), imminent threat (sudden appearance of human/dog), probable
satiation or other indecipherable reasons (for e.g., young juveniles can
indiscriminately pluck coconuts when learning the technique of ‘plucking and
dislodging coconuts’). We expressed
consumption and abandonment of coconuts in two different units across three
temporal scales, 1) as proportion in an annual coconut consumption cycle, 2) as
per capita mean in a month, and
3) as per capita mean throughout
the study. Similarly, coconuts used by
BW was expressed in two ways to reflect 1) overall share of different phenophases of coconuts and 2) proportionate share of
different phenophases of coconuts within demographic
classes. We compared (1) proportion data
using Chi-squared test of multiple proportion and (2) per capita figures across demographic classes, months,
annual feeding cycles and groups using parametric/non-parametric comparison of
means/ranks between two (e.g., t-test, Mann-Whitney U test) or more groups
(e.g., ANOVA, Kruskal-Wallis test). All
statistical analyses in this section were carried out using GraphPad Prism
v.8.3.1 (GraphPad Software 2020).
To test seasonality of coconut use by the demographic classes, we fitted
monthly per capita figures with
the standard equation for seasonality y = α + βsin(2πt) + γcos(2πt)
+ ε. In order to test the hypothesis
that (1) males have an overall greater consumption of processed coconuts than
females and (2) that late juveniles disproportionately determined use of
processed coconuts, we used a mixed effects modeling
approach using maximum likelihood estimation with Laplace approximation. We used coconuts consumed by a demographic
class (during a sampling day) as the dependent variable, month of sampling as
the random factor and group identity, age-class (computed monthly) and sex as
the fixed factors. To control for number
of individuals in a given demographic class, we used an offset term, log (#of
individuals). As a result of the
versatile computing ability of the R Statistical Programming Language, we used RStudio v.1.3 (RStudio
Team 2020) for all statistical analyses discussed in this section.
Finally, we illustrated frequency of entry to coconut plantation across
the day and represented duration of time spent by a group on entry at a given
time slot as mean ±SD. To depict trends,
we used a fifth order polynomial equation.
We plotted frequency of entry to coconut palm plantation alongside
corresponding duration of time spent in the plantation by collating data from
across all sampling days. All graphical
illustrations were carried out in GraphPad Prism v.8.3.1 (GraphPad Software
2020).
Ethical note
The present study was exclusively observational and did not involve any
invasive or controlled experimentation.
Clearance for the observational protocol was received from the
Institutional Animal Ethics Committee of the University of Mysore and complied
with the Code of Best Practices for Field Primatology.
Results
We undertook a total of 75 and 134 successful field samplings during a
period of 12 and 20 months during which we recorded a cumulative of 746 and 7,382
processed coconuts, and 243 and 566 unprocessed coconuts in TR and in BW,
respectively. Since a considerable
proportion of data emerged from scanning as opposed to direct observations,
information on demographic identity of the processing individual could not be
established. Hence, the dataset used for
demographic comparisons comprised a slightly smaller subset. We found evidences for variation in the use
of coconuts across the gradients of age, sex, group, and month (see Das et al.
2020). We describe the results of this
study below.
Age-specific acquisition of processed and unprocessed coconuts by Temple
Run group in a single annual cycle and by Baywatch group in two consecutive
annual cycles
We found contrasting results in the demographic shares of processed
coconuts between TR and BW groups though coconuts left unprocessed by the two
groups showed similar trends. With an
aggregate EJ:LJ:AD ratio of 4:5.8:5.5 TR showed the following crude order of
coconuts processed, LJ>AD>EJ (=267.17, df=1, p<0.0001;
=119.03, df=1, p<0.0001; =1148.16, df=1, p<0.0001; Figure 2). On the contrary, with an aggregate EJ:LJ:AD
ratio of 8:6.6:14 and 11.3:10.2:14 during the first (AC-1) and the second
annual cycles (AC-2), respectively, BW exhibited the following order of
demographic classes in the number of coconuts processed, AD>LJ>EJ
(=408.73, df=1, p<0.0001; =287.28, df=1, p<0.0001; =1532.14, df=1,
p<0.0001: =84.67, df=1, p<0.0001;
=1128.32, df=1, p<0.0001; =2027.09, df=1, p<0.0001; see Figure 3). An indicator of resources un-utilized and
perceived crop depredation by coconut horticulturists, the number of coconuts
left unprocessed were also assessed in a similar manner. We found late juveniles to be the highest
contributors to unprocessed coconuts across both troops and across both annual
cycles (in BW) coherently followed by adults and early juveniles (=313.49, df=1, p<0.0001; =7.73, df=1,
p=0.02; =464.18, df=1, p<0.0001:
=34.09, df=1, p<0.0001; =148.70, df=1, p<0.0001; =34.06, df=1,
p<0.0001: =26.92, df=1, p<0.0001;
=178.25, df=1, p<0.0001; =53.25, df=1, p<0.0001).
In absolute terms, late juveniles in TR abandoned coconuts 5.6 times
more than adults and 17.5 times more than early juveniles. Late juveniles in BW discarded coconuts 1.69
times and 1.67 times more than adults in the first and the second annual
cycles, respectively, and 4 times and 6.8 times more than early juveniles in
the first and the second annual cycles, respectively. Sex-specific shares of coconut consumption
across each demographic class are also presented in Figure 2 and in Figure 3.
Use of different phenophases of processed
coconuts by Baywatch group expressed as overall proportions and as demographic
class-specific proportions across two annual cycles
A stable pattern was revealed in phenophases
of coconuts used across both annual cycles.
The order of phenophases use emerged to be the
following P1>P2>P3/P4>P5 (=497.92, df=3, p<0.0001;
=1684.94, df=3, p<0.0001) (Figure 4). Comparison of absolute figures of phenophases consumed in the first annual cycle revealed P1
to be consumed more than P2 by a factor of 1.8, more than P3 by a factor of 3.5
and finally, more than P5 by a factor of 21.8 times. Similar figures in the second annual cycle
differed by a small margin with P1 consumed 1.2 times more than P2, 2.67 times
more than P3/P4 and finally, 104.6 times more than P5. Depiction of stacked columns of coconuts
processed by the demographic-classes in the first graph of Figure 4 is for
visual illustration alone.
To determine choice of phenophase of processed
coconuts used by a demographic class, we plotted the second graph of Figure
4. Note that the proportions within each
age-class add to unity. Individuals in the age-class of 13–36 months, i.e.,
early juveniles fed a disproportionately low number of P2 (7.5%) and P3
coconuts (3.8%) relative to P1 (88.7%) coconuts with no representation of P5
coconuts. Early juveniles showed the following order of preference based on the
number of coconuts processed, P1>P2=P3/P4 (=153.63, df=3,
p<0.0001) and P1>P2>P3/P4 (=435.37, df=3,
p<0.0001) during the first and the second annual cycles,
respectively. Late juveniles of the next
age category fed on all classes of phenophase and had
the following sequence of preference, P1>P2>P3/P4>P5(=287.80, df=3, p<0.0001; =823.60, df=3,
p<0.0001) across both annual cycles.
Proportion of P1 and P5 coconuts processed by late juveniles decreased
from 62.8% to 51.6% and from 1.7% to 0.4%, respectively whereas proportion of
P2 and P3/P4 processed coconuts increased from 26.4% to 32.8% and from 9.1% to
15.3%, respectively. Finally, adults
exhibited slight variability in their choice of coconuts across the two annual
cycles displaying the following order of preference, P1=P2>P3/P4>P5 (=167.68,
df=3, p<0.0001) in the first feeding cycle
and the order, P2<P1>P3/P4>P5 (=824.94, df=3,
p<0.0001) in the second feeding cycle (Figure 4). Corresponding alterations in the
proportionate consumption of different phenophases of
coconuts between the two annual cycles also became apparent, for example
increase in use of P1, P3/P4 and P5 coconuts from 35.1% to 41.1%, from 19.5% to
20.4% and from 0.5% to 3.3%, respectively and decline in the use of P2 coconuts
from 44.9% in the first annual cycle to 35.1% in the second annual cycle.
Abandonment of different phenophases of
coconuts (unprocessed) by Baywatch group expressed as overall proportions and
as demographic class-specific proportions
We found all phenophases of coconuts
represented in the unprocessed category of coconuts. We found that the phenophase(s)
that is/are processed the most is/are also the one(s) that is/are left
unprocessed the most; we found P1(50%) to be the highest unprocessed coconut in
the first annual cycle (=36.86, df=3, p<0.0001)
whereas both, P1(42.4%) and P2(30.9%) emerged as the phenophases
with the highest unprocessed coconut in the second annual cycle (=95.59, df=3, p<0.0001; =6.75, df=1,
p=0.08). The order of the
remaining phenophases did not show any consistent
pattern across the annual cycles. In the
first annual cycle, the proportion of P3/P4(26.5%) coconuts left unprocessed
was higher than P5 (7.4%) (=9.46, df=1, p<0.0001)
but equivalent to P2(16.2%) (=2.18, df=1, p=0.54)
whereas the proportion of P2 coconuts left unprocessed was comparable to P5
(=2.60, df=1, p=0.46). In the next annual cycle, we obtained the
following order of phenophases, P3/P4(22.03%) >
P5(4.66%) coconuts (=32.94, df=1, p<0.0001)
(Figure 5).
As opposed to processed coconuts, all the phenophases
were represented in unprocessed coconuts across all the demographic
classes. On analyzing
the proportion of phenophases of coconuts left
unprocessed by individual demographic classes across the annual cycles, we
found that the highest processed phenophase emerged
as the highest unprocessed coconut in the case of early juveniles (=60.80, df=3, p<0.0001) in the second annual cycle
alone. In the remaining demographic
classes however, no single phenophase of unprocessed
coconuts emerged as the single highest.
For instance, all phenophases were equally represented
among early juveniles in the first annual cycle (=8.00, df=3,
p=0.046); P1 and P3 were comparable in the first annual cycle (=7.06, df=1, p=0.07), and P1 and P2 were comparable in the
second annual cycle (=5.72, df=1, p=0.13)
among late juveniles and almost all phenophases were
equivalently left unprocessed by adults in both annual cycles (=5.78, df=3, p=0.12; =17.20, df=3,
p=0.0006). It is interesting to
note that P5 coconuts occurred in noticeable proportions (EJAC-1=12.5%,
LJAC-1=5.1%, ADAC-1=9.5%; EJAC-2=0%, LJAC-2=2.9%,
ADAC-2=8.8%) across all demographic classes in both annual cycles
except in the case of early juveniles in the second annual cycle.
Age-class specific monthly use of processed coconuts and abandonment of
unprocessed coconuts by Temple Run group
Over and above annual trends, we were interested in monthly patterns of
coconut use and coconut abandonment by demographic classes of the two groups
while controlling for class size, i.e., number of individuals in a demographic
class leading to the computation of per capita figures. We present the results of the two groups, TR
and BW, in separate sections followed by comparisons of the two groups in the
final section. All comparisons use per
capita values.
Considering a single annual cycle, while early juveniles and adults in
TR showed a stable use of processed coconuts across months (=24.22, p=0.01,
no difference between months on Dunn’s correction for multiple comparison;
=8.95, p=0.63), late juveniles showed a minor inter-month difference
(=2.17, p=0.03; µSep-2019>µMar-2019) (Figure
6). As a result of an almost constant
use of processed coconut across months and perhaps, lack of greater temporal
coverage, no seasonality was observed in the use of processed coconuts by any
of the demographic classes. At the level
of individual month, we found near-consistent difference between early
juveniles and late juveniles but no difference between late juveniles and
adults (Figure 6). In contrast, pooling
the data through the entire annual cycle showed a distinct demographic pattern
with LJ>AD>EJ (=112.50, p<0.0001) (Figure 6 inset). Similar to the analyses of processed
coconuts, we found no variation in the number of coconuts left unprocessed by
the demographic classes across months (=9.99, p=0.53; =21.66, p=0.03,
no difference between months on Dunn’s correction for multiple comparison;
=19.18, p=0.06; Figure 6) though across five of the 12 months, there
were minor differences across age-classes within a month in which late
juveniles emerged as the highest contributor to per capita
abandonment of coconuts. Consistent with our hypothesis, the following
order of coconuts left unprocessed emerged when data for the entire study were
pooled together, LJ>AD=EJ (=68.62, p<0.0001; ΣRankAD-ΣRankEJ=16.5,
p=0.053) (Figure 6 inset). Expectedly,
results from generalized linear mixed modeling
approach to determine relative influence of the three demographic classes on
use of processed and desertion of unprocessed coconuts showed that late
juveniles exerted greatest influence followed by adults and early juveniles
after controlling for variations due to month and number of individuals within
a demographic class (Table 1).
Age-class specific monthly use of processed and abandonment of
unprocessed coconuts by Baywatch
On comparing individual demographic classes in their use of processed
coconuts across months, we found both, early juveniles (=66.62, p<0.0001)
and adults to have unequal use (=5.99, p<0.0001) though late
juveniles showed a constant consumption pattern (=41.98, p=0.002; No
difference between months on Dunn’s correction for multiple comparison) across
the 20 months of the study. As a
consequence, no seasonality in the processing of coconuts was revealed in any
demographic class. Next, we attempted to
contrast the demographic classes at the level of individual months. The differences among the age-classes
appeared to be more subtle than TR since 11 out of the 20 months did not record
any difference among the age-classes.
Even among months (NEJ=7) where difference among age-classes
were retrieved, early juveniles emerged to have the lowest per capita
coconut processing value. Pooling data
from the 20 months of observation, we firmly established early juveniles to
have the lowest use of processed coconuts (µEJ±SD=1.12±0.95) but
late juveniles (µLJ±SD=2.24±1.41) and adults (µAD±SD=2.45±1.58)
had almost equal use of processed coconuts leading to the following order of
consumption, LJ~AD>EJ (=107.10, p<0.0001; ΣRankAD-ΣRankLJ=33.0,
p=0.054). A distinct difference among
the three demographic classes in BW was revealed with generalized linear mixed
model wherein, late juveniles emerged to exert greater influence on overall use
of processed coconuts than the remaining age classes (Table 1).
We analyzed data on unprocessed coconuts by
demographic classes in a manner similar to processed coconuts. Late juveniles did not vary in abandonment of
unprocessed coconuts across months (=48.59, p=0.0002; No difference
between months on Dunn’s correction for multiple comparison) though early
juveniles (=45.58, p=0.0006) and adults did (=58.54, p<0.0001)
(Figure 7). When demographic classes
were compared during each individual month, we found no difference among
age-classes during 11 months; at least one difference in paired comparison of
age-classes in seven months and just two months during which late juveniles
superseded both age-classes (March 2018 and December 2018). Considering overall aggregate figures of
unprocessed coconuts discarded by each demographic class, our result matched
the trend obtained for processed coconuts, AD=LJ>EJ (=107.10, p<0.0001;
ΣRankAD-ΣRankLJ=7.0, p>0.99)
(Figure 7 inset). Although, macaques of
both troops avoided heavily defended regions of palm plantations, we did not
find any difference in the number of coconuts left unprocessed among high (HR),
moderate (MR) and low risk (LR) areas of coconut plantations(=2.51, p=0.54; =3.21, p=0.33; =0.01, p=0.99; =4.15, p=0.13).
The results of the generalized linear mixed effects model considering processed
coconuts from both groups revealed that males superseded females by a mean
difference of 0.22 in per capita coconut consumption after adjusting for
age-class. We also found late juveniles
to have an overall highest mean per capita use of processed coconuts followed by adults
and early juveniles (Table 1) despite late juveniles in BW consuming an
equivalent number of coconuts as adults in BW.
Among other comparisons, BW and each representative age-class of BW had
higher mean per capita use of processed coconuts than TR and its
corresponding age-classes (Table 1). In
contrast, when unprocessed coconuts from both groups were pooled, BW and TR
were comparable along with early juveniles of both the troops (Table 2). Comparing age-classes of the two groups in
their abandonment of unprocessed coconuts, we found that adults of BW
superseded adults of TR and conversely, late juveniles of TR surpassed late
juveniles of BW. In addition, an overall
age-class related differences persisted where the mean difference in per
capita abandonment of coconut processing between LJ and AD, and between EJ
and AD were 1.15 and 0.58, respectively (Table 2).
Hourly pattern of entry into coconut plantations and corresponding
duration of foray by Temple Run and Baywatch groups
Temple Run exhibited a low but an approximately uniform frequency of
entry (~2–3 entries every 10 minutes) into plantations throughout the day with
minor peaks appearing at ~06.30h and at ~16.45h. An analogous trend was
obtained in pattern of duration of time spent which rose from an average of 0
min at 05.00h to 4 min at 06.00h and remained at an average of 10 min till
~17.00h before declining to 0 min.
Unlike the trends of foray duration obtained in TR, BW began their entry
into plantation slightly early and at a frequency of 16 (if the initial
frequency of 2 at 05.00h is ignored) at 05.10h after which, the trend of entry
declined sharply till 07.50h to null.
Frequency of entry gradually picked up to 6 till 14.50h and gradually
fell to null again at ~17.30h. Although
similar in form but widely different in magnitude, BW spent more time in
coconut plantations between roughly 09.30–13.30 h with peak average duration
reaching 250 minutes (as estimated from the polynomial trend line). Average duration of foray into plantation at
the tail of the trend line showed an asymptotic relationship with the straight
line slope of one (θ=45°) due to the constraint of activity period of the
species.
Discussion
In the first part of our article (Das et al. 2020), we applied the HIREC
framework to study differential use of coconut palms and palm plantations by a
wide variety of groups of long-tailed macaques differing in their exposure to
coconut and thus, to agricultural landscapes, habitat alterations and to biotic
threats from crop defending dogs/humans.
In the present article, we extended the HIREC framework to study the
response of demographic classes and decompose the use and abandonment of
coconuts. We found late juveniles to
consume and abandon the highest number of coconuts across annual cycles. On overall comparison by pooling data from
both groups, males emerged to use a higher share of processed coconuts than
females though sex-difference reversed in the late juvenile age class. Due to a skewed sex ratio (TR=1:4, BW=2:14)
in the adults, sex differences in the use of processed coconuts should be
interpreted with caution. Early juveniles constrained by their physiology and
limited processing repertoire almost exclusively used P1 coconuts though with
maturity, individuals used higher phenophases of
coconuts. Adult individuals were found
to equalize their choice of coconuts across phenophases
than late juveniles. The MVT construct
predicts patch-usage and patch-leaving decisions (e.g., Wajnberg
et al. 2000). Similar to results
obtained in the first part of the article (Das et al. 2020), entry into coconut
plantations were slightly more frequent during dawn and dusk though usage of
plantation remained stable through the day.
Temple Run showed a uniform frequency of use of coconut
clusters/plantations through the day but BW showed temporal modulations. The
corresponding profile of the distribution of ‘duration of foray’ between the
two groups, however, was highly variable.
The two groups also differed with regard to the spatial proximities of
their respective sleeping sites to nearest coconut plantations. Temple Run consistently chose sites that were
slightly aloof from human habitations and hence, distant from palm plantations. On the other hand, Baywatch had six of a
potential eight sleeping sites within 100m from palm plantations with three of
them located inside less-disturbed (i.e., low risk) regions of the
plantations. As a result of a relatively
wider (though sparse) distribution of coconut palm occurring within the range
of TR, they accessed coconut throughout the day in contrast to BW.
Age-specific use of processed coconuts and abandonment of unprocessed
coconuts by Temple Run group in a single annual cycle and by Baywatch group in
two consecutive annual cycles
The pattern of age-class governed unprocessed coconuts mirrored the
trend of processed coconuts. Dynamics of
the un-standardized age-class recorded in the dataset of BW, however, showed
less consumption of coconuts by late juveniles than adults due to a much
smaller count of late juveniles and as a result of stable demographic
structure, the trend remained constant over annual cycle. The disproportionately high records of
unprocessed coconuts abandoned by late juveniles relative to other age-classes
reflects indiscriminate and perhaps, naïve acquisition/plucking of coconuts
since a large subset is often unsuitable for consumption with coconut
abandonment, emerging as a byproduct of pedagogic
explorations of coconut. As is apparent, such explorative tendencies are
limited to young juveniles, since they are
physiologically/cognitively/mechanically constrained but not in adults as they
have the requisite cognitive and motor skills to harvest desired coconuts. In support of age-related proficiency of food
processing, description of cashew (Anacardium
spp.) processing by Wild Bearded Capuchins Sapajus
libidinosus also found age to be a strong
predictor of success in opening fresh and dry forms of the nut (Visalberghi et al. 2016).
Use of different phenophases of processed
coconuts by Baywatch group expressed as overall proportions and as demographic
class-specific proportions across two annual cycles
The frequency of use of phenophases of
processed coconuts followed the developmental order of phenophases
with the most immature stage(s) of coconut being used by all age-classes over
all the subsequent phases of coconut.
The share of P2 and P3/P4 phenophases in the
diet of early juveniles were meager and records were
made only from the oldest individuals in the category. Conversely, higher age-classes had greater
representations in mature phenophases of coconuts
with adults displaying skilled use of tougher/harder coconuts than late
juveniles. A study by Schaik & Noordwijk (1985) on Sumatran Long-tailed Macaques also
found adult males to select native wild fruits with hard rinds relative to
juveniles of <2 yrs. In contrast, Visalberghi et al. (2016) found that adult females process
a higher number of both dry and fresh cashews.
Analyses of the relative use of different phenophases
of coconuts within individual age-category clearly expounded age-related
patterns of resource use which denote a strong ontogenetic effect on extractive
foraging of coconuts. Similarly, balance
and optimality in choice of phenophases also seem to
be achieved at adulthood.
Abandonment of different phenophases of
coconuts (unprocessed) by Baywatch group expressed as overall proportions and
as demographic class-specific proportions
The trend of age-related unprocessed coconuts was incoherent across
annual feeding cycles, however, age-class with the highest explorative
tendencies was found responsible for the highest number of unprocessed
coconuts. It is intriguing to note that
despite being incapable to process P4 and P5 coconuts, early juveniles
proactively made efforts to dislodge and dehusk these
coconuts. For the remaining age-classes,
incidences of unprocessed coconuts were almost uniformly distributed across the
phenophases with P1-P3 showing highest incidences
among late juveniles in the first annual cycle and P1-P2 occurring in higher
numbers in the second annual cycle.
Curiously, despite their proficiencies in coconut processing, even
adults showed a substantial abandonment of P5 coconuts.
Age-class specific per capita use of processed and abandonment of
unprocessed coconuts by Temple Run and Baywatch groups across months
Representation of the monthly use of processed coconuts and abandonment
of unprocessed coconuts by age-classes distinctly identified late juveniles to
supersede the remaining age-classes though a slight difference was noted in the
month with the highest average per capita use/un-use (September in TR
and August in BW; similar to overall coconut consumption in Das et al.
(2020)). Analyses of pooled data from
the entire study period in the two troops reaffirmed the distinction of late
juveniles in TR though late juveniles were marginally comparable to adult
females in use of processed coconuts.
The difference in the results of the two groups is attributable to the
disparity in their food habits. As
opposed to the natural diet of BW, TR has a considerable dependence on
human-cultivated and artificially manufactured food items (Das et al. 2020). Late juveniles displayed adult-like use of
coconuts and developed commensurate sensorimotor and cognitive skills requisite
for extractive foraging, a direct evidence of high dietary dependence on the
drupe. Similar to Long-tailed Macaque,
Wild Bearded Capuchins were also found to show age-specific hierarchy in
average per capita processing of fresh cashew nuts. Adults and late juveniles processed equal
number of dry cashew nuts on an average (Visalberghi
et al. 2016). Remarkably, the
seasonality in the overall use of processed coconuts noted in Das et al. (2020)
failed to prevail when consumption was decomposed into age-classes. The absence of seasonality in coconut-use by
age-classes prompts us to speculate that as a consequence of coconut
scavenging, i.e., feeding kernel and/or drinking water from an already
processed coconut, cumulative harvest can satiate the entire group. Males consumed higher coconuts in all
age-classes, except in late juvenile stage possibly, as a result of dimorphism
in body sizes, which either, indicates non-coconut resource use by females or
that body-size maintenance by adult males trump energy requirements of
reproduction in females. Consistent with
the trend of crude comparisons of abandoned unprocessed coconuts by
age-classes, late juveniles had the highest overall per capita contributions to unprocessed coconuts,
especially in TR. Despite lower use and
hence, lower dependence on coconuts by late juveniles in TR relative to their
counterparts in BW, late juveniles in TR showed significantly higher
abandonment of coconuts indicating inefficacious handling and/or selection of
coconuts, an indication of suboptimal coconut foraging/processing
strategy. It is also to be noted that
coconuts left unprocessed can often be processed by the same or a different
individual and hence, is not a veritable index of crop loss.
Hourly pattern of entry into coconut plantations and corresponding
duration of foray by Temple Run and Baywatch groups
Temporal profile of entries to plantations and duration of forays were
largely modulated by the spatial distribution of coconut palms within the range
of groups. BW appeared to prefer
coconuts as their first choice of food at the beginning of the day,
strategically choosing sleeping sites that were either adjacent to or inside
palm plantations. Correspondingly, the
distribution of time spent in the morning during the first phase foray was very
high and coincided with the first foraging bout. Forays into plantations later in the morning
(i.e., after 07.20h) tended to be shorter than forays undertaken earlier (i.e.,
around 05.20h), possibly because sources of hydration and/or food has been
accessed. Subsequent use of plantations
through the day remained relatively low gradually increasing after 12.30h and
peaking at 15.30h, which corresponds to evening bouts of foraging. As a result of the edge distribution of the
plantations within the home range of the group, duration of time spent in
mid-afternoon tended to be longer as suitable foraging patches were distantly
located from the range edge. Patch-exit
decisions by BW were sporadically coerced by threats from humans/dogs guarding
the plantations, creating a landscape of fear that groups responded to even in
the absence of threat (Lindshield 2014; Gallagher et
al. 2017). For instance, arboreal paths
were preferred to enter risky areas of the plantation and nervous terrestrial
locomotion were noted among the most vulnerable members of the group as is
often reported in crop-foraging populations of non-human primates (e.g.,
Long-tailed macaques, Riley & Priston 2010;
Chimpanzee, Krief et al. 2014). The second group, TR on the other hand had
access to relatively uniform distribution of coconut palms/clusters throughout
its home range and therefore, entered/exited clusters regularly throughout the
day spending almost equivalent duration throughout the day. Hostilities, however, did not have any effect
on proportion of coconuts abandoned in riskier human areas. The lack of sharp peaks in the temporal
distribution of plantation entry could also be a strategy in response to
anthropogenic hostilities faced there.
By flattening the temporal curve of plantation entry through the day,
the probability of occurring in plantations through the day although low,
becomes finite and hence, possibility of facing resistance becomes low. Therefore, alongside physiological (like
hunger and thirst) and resource-based (like abundance, distribution and
nutrition) factors, spatiotemporal pattern of threat from human/dog modulates
resource patch usage. Applying the
HIREC-MVT construct, we infer that coconuts are highly energetic sources of
food and nutrition for the species and are lucrative enough to risk entry into
moderately-defended portions of plantations.
Selective pressures of this human-macaque interface especially, for an
edge population has also prompted the development of surreptitious foraging
tactics in both the groups, exemplified by suppression of vocal communication,
heightened vigilance, spurts of rapid movements and controlled motor actions to
reduce noise.
To summarize, extractive foraging of an embedded and heavily-defended
cultivar, like coconut have challenged macaques in many ways. For example, the embedded/encased nature of
the fruit permits early juveniles to exploit only tender phenophases
of coconuts. Adults face similar hurdles
with mature stages of coconuts and hence have a balanced choice of phenophases that optimizes net benefits. Even context-specific choice of phenophases by adults though not explored in this article
is suspected, which could further elucidate cognitive proficiencies of adults
in determining suitability of coconut.
It is remarkable to note that description of the use of a single dietary
resource generated the order of consumption precisely as predicted by the
theory of life-history strategies. A
second class of challenge emanates from coconut foraging from highly defended
plantations, which is studied by describing temporal strategies of plantation
entry and plantation use. The groups
were found to employ deceptive strategies suited to minimize detection by
maintaining the probability of entry to plantations at a non-zero level through
the day and by adopting covert communications and clandestine movements inside
plantations, a subject matter that we will further explore.
Table 1. Results of the generalized linear mixed effects model with
number of processed coconuts consumed as the dependent variable, month as the
random factor and age-class, group identity and sex as the fixed factors. To control for number of individuals in an age-class,
we have used an offset term, log (number of individuals). Model fitting has used maximum likelihood
method along with Laplace approximation.
Coefficient |
Estimate (β) |
SE |
Z |
p |
Processed coconuts~ Age class * Sex + Age Class*Group + (1 |Month) |
||||
Intercept |
0.268 |
0.097 |
2.762 |
0.0058 |
Early juvenile |
-1.739 |
0.097 |
-17.89 |
<0.0001 |
Late juvenile |
0.221 |
0.042 |
5.26 |
<0.0001 |
Male |
0.218 |
0.050 |
4.40 |
<0.0001 |
Temple Run |
-1.243 |
0.075 |
-16.54 |
<0.0001 |
Early juvenile: Male |
0.520 |
0.118 |
4.40 |
<0.0001 |
Late juvenile: Male |
-0.198 |
0.067 |
-2.94 |
0.0032 |
Early juvenile: Temple Run |
-0.516 |
0.217 |
-2.375 |
0.0175 |
Late juvenile: Temple Run |
0.752 |
0.088 |
8.20 |
<0.0001 |
*Adult (Age class), Female (Sex class) and Baywatch (Group) have been
used as reference categories
Table 2. Results of the generalized linear mixed effects model with
number of unprocessed coconuts consumed as the dependent variable, month as the
random factor and age-class, group identity and sex as the fixed factors. To control for number of individuals in an
age-class, we have used an offset term, log (number of individuals). Model fitting has used maximum likelihood
method along with Laplace approximation.
Coefficient |
Estimate (β) |
SE |
Z |
p |
Unprocessed coconuts ~ Age Class*Group + (1 |Month) |
||||
Intercept |
-2.293 |
0.113 |
-20.259 |
<0.0001 |
Early juvenile |
-0.577 |
0.137 |
-4.206 |
<0.001 |
Late juvenile |
1.152 |
0.090 |
12.768 |
<0.0001 |
Temple Run |
-0.338 |
0.203 |
-1.662 |
0.0966 |
Early juvenile: Temple Run |
0.070 |
0.369 |
0.189 |
0.850 |
Late juvenile: Temple Run |
0.767 |
0.217 |
3.528 |
<0.001 |
*Adult (Age class) and Baywatch (Group) have been used as reference
categories
For figures
and images - - click here
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